Biological Report 82 (11.60)June 1986

TR EL=82-4

Species Profiles: Life Histories andEnvironmental Requirements of Coastal Fishesand Invertebrates (Pacific Southwest)

STEELHEAD

Coastal Ecology GroupFish and Wildlife Service Waterways Experiment StationU.S. Department of the Interior U.S. Army Corps of Engineers

Biological Report 82(11.60)TR EL-82-4June 1986

Species Profiles: Life Histories and Environmental Requirementsof Coastal Fishes and Invertebrates (Pacific Southwest)

STEELHEAD

by

Roger A. BarnhartCalifornia Cooperative Fishery Research Unit

Humboldt State UniversityArcata, CA 95521

Project OfficerJohn Parsons

National Coastal Ecosystems TeamU.S. Fish and Wildlife Service

1010 Gause BoulevardSlidell, LA 70458

Performed for

Coastal Ecology GroupWaterways Experiment StationU.S. Army Corps of Engineers

Vicksburg, MS 39180

and

National Coastal Ecosystems TeamResearch and DevelopmentFish and Wildlife Service

U.S. Department of the InteriorWashington, DC 20240

CONVERSION TABLE

Multiply By To Obtain

millimeters (mm) 0.03937 inchescentimeters (cm) 0.3937 inchesmeters (m) 3.281 feetkil ometers (km) 0.6214 miles

square meters (m2) square kihectares

ometers.(kmL)ha)

liters (1) 0.2642cubic meters (m3)

'gallons35.31 cubic feet

cubic meters 0.0008110 acre-feet

milligrams (mg) 0.00003527 ouncesgrams (g) 0.03527 ounceskilograms (kg) 2.205 poundsmetric tons (t) 2205.0 poundsmetric tons 1.102 short tonskilocalories (kcal) 3.968 British thermal units

Celsius degrees 1.8("C) + 32 Fahrenheit degrees

inches 25.40 millimetersinches 2.54 centimetersfeet (ft) 0.3048 metersfathoms 1.829 metersmiles (mi) 1.609 kilometersnautical miles (nmi) 1.852 kilometers

square feet (ft2)acressquare miles (mi2)

0.0929 square meters0.4047 hectares2.590 square kilometers

gallons (gal) 3.785 literscubic feet (ft3) 0.02831 cubic metersacre-feet 1233.0 cubic meters

ounces (oz)pounds (lb)short tons (ton)

28.35 grams0.4536 kilograms0.9072 metric tons

British thermal units (Btu) 0.2520 kilocalories

Fahrenheit degrees 0.5556("F - 32) Celsius degrees

iv

Metric to U.S. Customary

10.76 square feet0.3861 square iniles2.471 acres

U.S. Customary to Metric

.

I’

CONTENTS

Page

PREFACE ...................................................................... iiiCONVERSION TABLE .............................................................. ivACKNOWLEDGMENTS ............................................................... vi

NOMENCLATURE/TAXONOMY/RANGE .................................................. 1MORPHOLOGY/IDENTIFICATION AIDS ...............................................1REASON FOR INCLUSION IN SERIES ............................................... 4LIFE HISTORY ................................................................. 4THE FISHERY .................................................................. 6ECOLOGICAL ROLE .............................................................. 8ENVIRONMENTAL REQUIREMENTS ................................................... 9

Temperature ................................................................. 9Dissolved Oxygen ........................................................... 10Depth ...................................................................... 11Water Movement ............................................................. 11Sediment ................................................................... 12

LITERATURE CITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

V

ACKNOWLEDGMENTS

I thank Dennis P. Lee, California Department of Fish and Game, and Terry D.Roelofs, Humboldt State University, for reviewing the manuscript; Thomas H.Hassler, California Cooperative Fishery Research Unit, for acting as the liaisonwith the National Coastal Ecosystems Team; and Delores Neher for greatlyfacilitating the preparation of this report.

vi

Figure 1. Steelhead.

Ld STEELHEAD

NOMENCLATURE/TAXONOMY/RANGE

Scientific name . . . . . . . . Salmogairdneri Richardson

Preferred common name . . . Steelhead(Figure 1)

Other common names steelheadtrout, steelie, half-pounder, iron-head

Class . . . . . . . . . . OsteichthyesOrder . . . . . . . . . SalmoniformesFamily . . . . . . . . . . Salmonidae

Geographic range: At sea, fromnorthern Baja California to theBering Sea and Japan. In thePacific Southwest, steelhead entercoastal streams from the northernCalifornia border south to theVentura River (Figure 2). Adultsteelhead have been reported in theSanta Clara and Santa MargaritaRivers in years when winter runoffwas high enough to allow upstream

migration (Swift 1975). Early inthis century steelhead distributionin the Pacific Southwest was morewidespread (Figure 3). Steelheadrun up the Santa Domingo River in northern Baja California during highrunoff in winter (Needham and Gard1959).

MORPHOLOGY/IDENTIFICATION AIDS

The following descriptions weretaken from McConnell and Snyder (1972)and Fry (1973).

At sea, steelhead are steel blueabove and bright silver on the sidesand belly. Sharply defined blackspots are located on back, head,sides, and dorsal and caudal fins.The spots are small and vary innumber. After entering freshwater,steelhead gradually take on the

1

124O ILOO

OREGON42O-

NEVADA

CALIFORNIA

PACIFIC

OCEAN

1

IN

0 50 100mi, 1 Ir I I0 100 200km ANGELES

- 33O & Terminal Dam

Figure 2. Distribution of steelhead (shaded areas) in Pacific Southwest regionstreams in 1980's (California Department of Fish and Game, pers. comm.; Moore1980).

2

PACIFIC OCEAN

1 N

0 50

0 100

CALIFORNIA

IOOmi

200km

Figure 3.Distribution of steelhead (shaded areas) in the Pacific Southwest region in 1900 (National Council on.Gene Resources 1982).

3

appearance of stream rainbow trout;the back becomes olive green and thesides and belly less silvery.Maturing adults usually develop abroad pink stripe along the lateralline and pink coloration on theoperculum. Steelhead lack the redstreaks beneath the jaw whichcharacterize the cutthroat trout(Salmo clarki). Juvenile steelhead in- -streams cannot be distinguished fromjuvenile resident rainbow trout.

Anal rays 9-12, rarely 13; teethon tip and shaft of vomer and ontongue; gill rakers on lower limb offirst arch range from 17-21; scales infirst row above lateral line 115-180;caudal fin shallowly forked; maxillarydoes not reach past the posteriormargin of eye; juvenile parr markssmall and oval to nearly round.

Generally adult steelhead in thePacific Southwest region weigh lessthan 4.5 kg but some exceed 11 kg.

REASON FOR INCLUSION IN SERIES

The steelhead is the mostwidespread anadromous sport fish inthe Pacific Southwest region, but itsabundance has declined since the1950's because of increased sportfishing intensity and damage to fishhabitat through construction of dams,water diversion, road construction,and improper land managementpractices. In some rivers a steelheadfishery would be nonexistent withouthatchery stocks.

I n C a l i f o r n i a , attempts arebeing made to protect wild stocks ofsteelhead to maintainspawning and

existingrearing habitat, to

restore or enhance degraded habitat,to use artificial propagation wisely,and to regulate the fishery to providequality angling (California Departmentof Fish and Game 1975).

LIFE HISTORY

Steelhead trout spend a portionof their life in the ocean where mostof their growth occurs and sexualmaturity is attained; then they enterfreshwater to spawn. Spawning gener-ally takes place from February to lateJune. The eggs are laid in pits(redds), dug in the gravel of thestream bottom by the female. Imme-diately after the eggs are laid andfertilized, they are covered withgravel by the female; the length oftheir stay in the gravel depends uponwater temperature, dissolved oxygenconcentration and substrate composi-tion. After the eggs hatch, the youngsteelhead gradually work their way tothe surface of the stream bed. Juve-niles usually spend a year or longerin freshwater (length of residence isdetermined by environmental andgenetic factors) and then descend tothe ocean.

The steelhead is a strain ofrainbow trout that has a strong urgeto migrate to the ocean; however, someindividuals may remain in a stream,mature, and even spawn without goingto sea (Shapovalov and Taft 1954).

The life history of the steel-head trout varies more than that ofany other anadromous fish regardingthe length of time spent at sea, thelength of time spent in freshwater,and the times of emigration from andimmigration to freshwater. Unlikesalmon, steelhead do not usually diejust after spawning.

Steelhead are classified intotwo races (Withler 1966; Smith 1960,1969; and Everest 1973): wintersteelhead that enter streams betweenNovember 1 and April 30, and summersteelhead that enter streams betweenMay 1 and October 30. Portions ofboth groups may enter freshwater inspring or fall and are then calledspring- or fall-run steelhead. Inlarge rivers, such as the Klamath andSacramento Rivers, steelhead may enter

4

during most of the year. Winter-runsteelhead usually enter freshwater asmaturing fish that spawn relativelysoon, whereas most summer steelheadenter as immature fish and do notmature and spawn until several monthslater. In the Pacific Southwest,summer steelhead are not abundant andthe runs in many streams consist ofless than 100 fish (Roelofs 1983).Degradation of habitat in inter-mittent streams and susceptibility toangling and predation probably accountfor the low numbers. The southernmostsummer steelhead population is in theMiddle Fork Eel River. This riversupports the largest run of these fishin the Pacific Southwest--up to 2,000fish in a good year.

An exception is the "half-pounder" summer steelhead (terminologyof Snyder 1925) which has a uniquelife history described by Kesner andBarnhart (1972) and Everest (1973).Half-pounder runs are confined to asmall geographic range encompassingabout 120 miles of the southern Oregonand northern California coasts,including the Rogue, Klamath, Mad andEel Rivers. These small immaturesteelhead (25-35 cm long) annuallyenter freshwater from late August toearly October, and are the basis ofimportant sport fisheries in theKlamath and Rogue systems. Half-pounders spend only a few months atsea before they return to freshwaterand, in contrast to mature steelhead,feed extensively in freshwater. Half-pounders that survive their firstupstream migration return to the oceanthe following spring and migrate backto freshwater as mature steelhead inthe summer and fall. Everest (1973)reported that half-pounders annuallycomprise about 65% of the summer-runof steelhead on the Rogue River andthat 97% of all adult summer steelheadfrom there make their first upstreammigration as half-pounders.

Sumner and winter steelhead donot interbreed; they are isolatedtemporally and spatially (Smith 1969;

Everest 1973). Sumner steelhead spawnin January and February, whereaswinter steelhead spawn in April andMay, and summer steelhead spawn insmaller streams or farther upstream.The sex ratio of steelhead troutimmigrants is about 1:1 (Shapovalovand Taft 1954; Kesner and Barnhart1972). Female steelhead containabout 2,000 eggs per kilogram of bodyweight (Moyle 1976).

Several researchers haveconcluded that the incidence of steel-head spawning more than once increasesfrom north to south (Bali 1959;Withler 1966; Sheppard 1972). Widevariations in the percent of repeatspawning can be due to geneticfactors, habitat quality, fishingintensity, and management practices.Fish that have spawned twice make up70% to 85% of repeat spawners, whereasthose that have spawned three timesmake up 10% to 25% of all repeatspawners (Forsgren 1979). The fewfish that have spawned four times arelikely to be females, which have ahigher survival rate than males duringand following spawning. Spawningmales usually each serve more than onefemale, remain in the stream longerthan the females (tagging studies byJones (1974) indicated nearly twoweeks longer), and are exposed to moreprolonged physical exertion than thefemales (Meigs and Pautzke 1941).

Steelhead spawn in cool, clear,well-oxygenated streams with suitabledepth, current velocity, and gravelsize (Reiser and Bjornn 1979). Mea-surements made over steelhead reddsshowed that steelhead spawn at depthsof 0.10-1.5 m, current velocities of23-155 cm/sec, and in gravel of 0.64-12.70 cm in diameter (Smith 1973;Hunter 1973; Bovee 1978; Wesche andRechard 1980). Intermittent streamsare often used by steelhead forspawning (Everest 1973; Kralik andSowerwine 1977; Carroll 1984). Mostof the fry produced emigrate to peren-nial streams soon after hatching.

5

The embryology of the steelheadis similar to that of other salmonids(Wales 1941). The number of daysrequired for steelhead eggs to hatchvaries from about 19 at an averagewater temperature of 15 'C to about 80days at an average of 5 'C. Steelheadfry usually emerge from the gravel 2to 3 weeks after hatching.

After emergence, steelhead fry(often in small schools) usually livein shallow water along the streambanks. As the fry grow older theschools break up and the individualfish establish territories which theydefend. Most steelhead in their firstyear of life tend to inhabit rifflesbut some of the larger fish inhabitpools or deeper faster runs. Mortal-ity is high during the first fewmonths after emergence and manyinvestigators have suggested that therelative size of the year class islargely determined at that time(Chapman 1966; McFadden 1969; Burns1971; Everest and Chapman 1972).

In recent years, habitat degra-dation has lowered the capacity ofmany streams to rear steelhead tosmolts . For example, excessive sedi-mentation has reduced food production,pool depth, and cover--all importantto juvenile steelhead survival.

Juvenile steelhead feed on awide variety of aquatic andterrestrial insects. Newly emergedfry sometimes are preyed upon by olderjuvenile steelhead. Young steelheadmoving about trying to find a suitableterritory are subject to the highestpredation (Shapovalov and Taft 1954;Chapman 1966).

Juvenile steelhead live infreshwater for from 1 to 4 years

'Smelt: Term applied to an anadromousjuvenile salmon‘id that is physiologi-cally prepared to adapt to a saltwaterexistence; steelhead smolts lose theirparr marks and become silvery.

(usually 2 years in the Pacific dSouthwest) before becoming smolts andmigrating to saltwater. Most of themigrants are 2 years old and 18.6 to21 cm long (fork length) in theKlamath River (Kesner and Barnhart19972), or 14 to 21 cm long in WaddellCreek (Shapovalov and Taft 1954). Thelarger the smolts, the better thesurvival. Most steelhead smolts enterthe sea in March and April.

Steelhead live 1 to 4 years inthe ocean (usually 1 or 2 years in thePacific Southwest). The length ofresidence in both freshwater and salt-water increases from south to north(Withler 1966). Steelhead growrapidly in the ocean and their sizeupon reaching maturity depends pri-marily on how long they have lived inthe ocean. In California, the averagelength of adult steelhead after 2years in the ocean is 58 cm (Withler1966). Immature steelhead trout (half-pounders) increase about 30 mm inlength each month from the time theyenter the ocean until they return tofreshwater (Kesner and Barnhart 1972).

The distribution or migration ofsteelhead in the ocean is not welldefined, particularly in the PacificSouthwest. As judged by tag returns,most steelhead tend to migrate northand south along the Continental Shelf.Steelhead stocks from the Klamath andRogue Rivers probably mix together ina nearshore ocean staging area alongthe northern California coast beforethey migrate upriver (Everest 1973).

THE FISHERY

Commercial fishing for steelheadhas been prohibited in the PacificSouthwest since 1924. The species nowis managed exclusively as a sportfishery. Most steelhead are caught inrivers rather than in the ocean. Thethree most important steelhead troutrivers are the Klamath and Eel Riversin northern California andSacramento River which empties

theinto

d

6

San Francisco Bay. The CaliforniaDepartment of Fish and Game (1965) inits Fish and Wildlife Plan estimatedspawning runs of 221,000 steelhead inthe Klamath River and 82,000 steelheadin the Eel River. The Klamath Riverstill has good runs of nativesteelhead, perhaps approaching 200,000fish in good years (Barnhart 1975).In the upper Sacramento River from1953 to 1959, the average run of adultsteelhead was 20,590 and consisted ofboth natural and hatchery-producedfish (Hallock et al. 1961).

In the California Fish and Wild-life Plan (1965), the State's annualspawning stock of steelhead was esti-mated at 603,000 fish. The Plan alsoreported that the State contained8,402 linear miles of steelhead habi-tat, 31% of which was available toanglers. It was estimated that steel-head anglers fish 304,000 angler-daysper year to harvest 122,000 fish(about 0.4 fish per angler-day). Many

iCalifornia streams support a substan-tial fishery for juvenile steelhead--fish that have not yet migrated to theocean. These 5- to 8-inch trout aretaken in large numbers, mostly duringthe so-called summer trout season.The fishing pressure for these trout(about 440,000 angler-days annually)exceeds that on adult steelhead (Cali-fornia Department of Fish and Game1965).

Although adult steelhead do notcommonly feed in freshwater, they arereadily taken by angling. Bait fish-ing is popular and effective. Salmoneggs or clusters of roe or nightcrawlers are drifted through riffles,pools, and especially where a tribu-tary enters a large stream. Spinnersand other artificial lures are alsoused. When streams are low and clear,artificial flies are sometimes effec-tive. Most steelhead fishing is donefrom the bank or by wading; driftingin boats is popular in larger streams.

The decline of wild, naturallyproduced steelhead in the spawning

7

runs in the Pacific Southwest regionsince the 1940's has required an in-crease in the stocking of hatcheryreared steelhead. A total of about 1.9million yearling steelhead for plant-ing are supplied annually by ColemanNational Fish Hatchery on BattleCreek, a tributary to the upper Sacra-mento River, the Nimbus Fish Hatcheryon the American River, the FeatherRiver Hatchery on the Feather River,and the Mokelumne River Hatchery onthe Mokelumne River. The goal is tomitigate the loss of steelhead troutin those rivers that has been causedby dams and water diversion.

The average weight of hatcheryyearlings at the time of planting(January through May) is about 51grams (9 fish per pound). At anexpected return of about 2% annually,planted steelhead contributed about38,000 adults to the run in theSacramento River (Hallock et al.1961). If steelhead were not stocked,the catch in the Sacramento Riverwould be much smaller. The CaliforniaDepartment of Fish and Game operatesfour hatcheries on the coast thatproduce about 1.5 million yearlingsteelhead annually. In addition,trout rearing projects sponsored bythe California Department of Fish andGame, private groups, and countiescontribute an average of 114,000yearlings annually (Boydstun 1977a).On the Klamath system, where two ofthe coast hatcheries are located,hatchery production contributes onlyan estimated 8% of the run of adults(Boydstun 1977b). California alsoreleases nearly 3 million steelheadfingerlings annually (Jensen 1971).

Fishery regulations are used tohelp protect declining steelheadstocks from excessive fishing. InCalifornia, the daily bag limit isthree adult fish. All streams tribu-tary to major coastal rivers and mostsmall coastal streams are closed toangling from mid-November to lateApril. Special regulations also aresometimes used. For example, a

section of the upper Middle Fork EelRiver has been permanently closed toangling to protect the adult summersteelhead trout that concentrate therein summer and fall.

The use of hatchery reared troutto boost populations is not entirelybeneficial. Larger runs attract morefishermen, and more fishermen furtherreduce the abundance of wild stocks.Also, the genetic mixing of hatcheryand wild stocks could decrease theadvantageous traits in wild stocks.The steelhead trout policy of theCalifornia Department of Fish and Gamelimits artificial propagation andstocking to reduce such interferencewith natural Salmonid stocks; suchmeasures are periodically reviewed toassess their effects on the wildstocks.

The environmental quality ofcoastal rivers largely governs thelevel of steelhead production. I n thePacific Southwest, attempts have beenmade to increase the protection ofexisting spawning and rearing habitatand to rehabilitate damaged habitatwhere feasible. California's steel-head trout policy is directed towardthree goals: (1) to protect andimprove steelhead trout habitat; (2)to develop plans and programs forassessment of habitat conditions andadverse impacts, land use planning,and acquisition of interests instreams threatened with adverse devel-opments; and (3) to study the effectsof habitat changes caused by over-grazing, gravel extraction, logging,road construction, urbanization, andwater development.

In 1982-84, California spent$900,000 annually on the restorationof salmon and steelhead habitat. ASalmon Restoration Project was devel-oped within the California Conserva-tion Corps in 1980, and by the end of1983, Corps enrollees removed 7,447cords of wood and debris from 233miles of streams used by anadromousfish (Kreb 1984). The Conservation

Corps also planted streamside vegeta-tion and constructed fish passageweirs on some streams. In 1981-1983,an additional $1 million was spentannually by California to enhancespawning gravels on the upper Sacra-mento River and the Shasta and KlamathRivers, and to construct Salmonidrearing ponds (Rawstron and Hashagen1984). Six Rivers National Forest innorthern California during the past 5years has carried out an extensiveprogram to restore or enhance thespawning and rearing habitats ofanadromous fish (Overton et al. 1981;Overton 1984).

ECOLOGICAL ROLE

In freshwater, young steelheadmay be sympatric with sculpins, resi-dent rainbow trout, coho salmon(Oncorhynchus kisutch), and in someinstances other salmonids. Data arelacking on the effects of interactionof juvenile steelhead and residentrainbow trout, largely due to thedifficulty in distinguishing betweenthe fish. In the Pacific Southwest,resident rainbow trout are common instreams above barriers to steelheadmigration and fry sometimes driftdownstream over the barriers. Bjornn(1978) reported that steelhead frytended to displace juvenile residenttrout in the Lemhi River, Idaho. Theecological requirements of the tworaces are similar.

Coho salmon and steelhead troutare similar in geographical distribu-tion, systematic characteristics,spawning locations, food habits, andthe length of time the young spend infreshwater (Milne 1948), althoughsteelhead normally remain infreshwater longer than 1 year.Although both species live in similarhabitat in their first year, Hartman(1965) reported that in spring andsummer, most steelhead trout live inriffles and most coho salmon live inpools. Several investigators havereported that spatial segregation is

1

J

3

also vertically stratified; cohosalmon live near the stream surfaceand steelhead near the bottom (Hartman

t1965; Peterson 1966; Edmundson et al.1968; Bustard and Narver 1975).

IAlthough coho salmon hatch earlier andconsequently are larger at first,steelhead fry grow so much faster thatby late summer the size difference issmall. Interspecific competition isprobably not serious because of theinitial difference in size, differ-ences in habitat preference, and thedifference in age at first emigrationto the sea (Fraser 1969). Much thesame relation was reported for juve-nile chinook salmon (Oncorhynchustshawytscha) and juvenile steelhead byChapman and Bjornn (1969).

Native and hatchery-rearedsteelhead exhibit some competition.Steelhead trout planted at the wrongtime or at the wrong size tend to stayin the stream longer than usual and

L

are more competitive with wild fish(Royal 1972). Heavy predation byhatchery-raised steelhead smolts onchinook salmon fry below ColemanNational Fish Hatchery on theSacramento River was reported byMenchen (1981). Wagner (1967) statedthat ideally the stream is to serveonly as a highway to the sea and notas a postliberation rearing area forhatchery products. Pollard and Bjornn(1973) reported that the stocking ofhatchery rainbow trout also caused alocalized temporary decrease in theabundance of juvenile steelhead.

In coastal streams, steelheadfry are preyed on by sculpins (Cottusspp. ) , larger steelhead, and rainbowtrout (Shapovalov and Taft 1954); bybirds such as the great blue heron(Ardea herodias), belted kinqfisher

-dippercommon

mergansers (Mergus merganser); bygarter snakes (Thamnophis spp.); and

by various mammals (Shapovalov andTaft 1954; Sheppard 1972; Cross 1975).In the ocean, steelhead are eaten by

fish and marine mammals but the extentand effect of predation are unknown.

In freshwater, steelhead feedprimarily on immature aquatic stagesof insects and secondarily on matureterrestrial insects. Ephemeroptera(mayflis), Diptera (true flies), andTrichoptera (caddisflies) are usuallythe most important taxa in the diet(Shapovalov and Taft 1954; Royal 1972;Fite 1973; Hiss 1984). Juvenilesteelhead are somewhat opportunistic,feeding on almost any available insect(Fite 1973). The size of theterritory for a single juvenile isdetermined largely by the availabilityof food and the size of the fish(Allen 1969). In the ocean, steelheadfeed on a variety of organismsincluding juvenile greenlings(Hexagrammos spp. ), squids, and amphpods (LeBrasseur 1966; Manzer 1968).

i-

ENVIRONMENTAL REQUIREMENTS

Temperature

Water temperature affects allmetabolic and reproductive activitiesof fish, including such criticalfunctions as growth, swimming, and theability to capture and assimilate food(Tebo 1974). Optimum temperaturerequirements may vary, depending onthe season and life stage. Aproductive steelhead stream shouldhave summer temperatures in the rangeof 10 to 15 ‘C and an upper limit of20 Oc. Steelhead have difficultyextracting oxygen from water attemperatures much over 21 Ocregardless of the amount of oxygenpresent (Hooper 1973). Bell (1973)listed the preferred temperatures ofyoung steelhead as 7.2 to 14.5 'C, theoptimum as about 10 'C, and the upperlethal limit as 23.9 OC. Bovee(1978), who developed a probability-of-use temperature curve for rearingwinter steelhead juveniles,wrote thatoptimum temperatures ranged from 0 to24 'C and peaked at 15 'C. In studiesof the smolting of cultured steelhead

in the spring, Wagner (1974) andKerstetter and Keeler (1976) reportedthat low temperatures extend thesmolting period and high temperaturesshorten it. They reported that smolt-ing ceased rather abruptly when watertemperatures increased to 14-18 'C.

During the spawning season asudden drop in water temperature maycause all salmonid spawning activityto cease (Reiser and Bjornn 1979).Reingold (1968) reported that watertemperatures of 2 to 10 'C impairedthe viability of eggs and delayed theripening of adult steelhead held for51 days in a hatchery pond. Bovee(1978) reported a spawning temperatureran e for winter steelhead of 4 to13 a C, and a peak of 8 'C. Bell(1973) indicated that steelheadspawning temperatures are generallyfrom 3.9 to 9.4 OC. The averagedevelopment time from fertilization tohatching lengthens considerably withdecreasing temperature; f8r

rainbowtrout it is 19 days at 15 C, 31 daysat 10 'C, and 80 days at 5 OC (Embody1934). Bovee (1978) gave anincubation temperature range forwinter steelhead of 0 to 24 'C and anoptimum temperature of about 10 'C.

When water temperatures fallbelow 4 'C in streams of the PacificNorthwest, juvenile steelhead becomeinactive and hide in available coveror in the substrate (Chapman andBjornn 1969; Bustard and Narver 1975).In California's coastal streams,juvenile steelhead remain active year-round and in one small coastal stream,young steelhead grew throughout thewinter (Reeves 1979).

The virulence of many fishdiseases and the toxicity of mostchemicals increase with increasingwaterRemovalresultstreamreduced1971).

temperatures (Lantz 1971).of riparian vegetation can

in marked increases in summertemperatures and sometimes inwinter temperatures (Brown

Dissolved Oxygen

Stream-dwelling salmonids requirehigh dissolved oxygen in both thewater column and intragravel waters.The swimming performance of juvenileand adult salmon was impaired when thedissolved oxygen concentration wasreduced below the air-saturation level(Davis et al. 1963). A sharp decreasein performance was noted at 6.5-7.0mg/l Reiser and Bjornn (1979) wrotethat the oxygen levels recommended forspawning anadromous fish (at least 80%of saturation, with temporary levelsno lower that 5.0 mg/l) should meetthe needs of migrating salmonids.

Intragravel dissolved oxygenconcentration is positively related tosurvival of salmonid embryos (Coble1961; Phillips and Campbell 1962;Silver et al. 1963). Silver et al.(1963), in a controlled laboratoryexperiment conducted at 9.5 OC, foundthat steelhead embryos hatchedsuccessfully at mean oxygen concen-trations as low as 2.6 mg/l but thattotal mortality occurred at a meanlevel of 1.6 mg/l. In a field experi-ment Phillips and Campbell (1962)noted that total mortality ofsteelhead embryos occurred at meanoxygen concentrations of 7.2 mg/l orless. Although intragravel oxygenlevels may appear adequate, the amountof oxygen actually reaching theembryos also depends on the intra-gravel water velocity (Wickett 1954).The amount of intragravel oxygenavailable to developing embryos issometimes reduced by the biochemicaloxygen demand of organic material inthe gravel bed (Ponce 1974). Even ifembryos hatch at low or moderatelyreduced oxygen levels, the incubationperiod is extended and the resultingfry are likely to be smaller andweaker than those reared at oxygenconcentrations close to saturation(Silver et al. 1963; Shumway et al.1964). Reiser and Bjornn (1979)concluded that although dissolvedoxygen concentrations required for

l

successful incubation depend on bothspecies and developmental stage,concentrations at or near saturationwith no temporary reductions below 5.0mg/l are generally required by anadro-mous salmonids.

In Salmonid nursery and rearingstreams dissolved oxygen concentra-tions of surface waters are normallynear saturation, except in smalltributaries with large amounts ofdebris from logging or other sourcesor in large, slow-moving streamsreceiving large amounts of municipalor industrial waste (Reiser and Bjornn1979). Salmonids function normally atdissolved oxygen concentrations of7.75 mg/l; exhibit various distresssymptoms at 6.00 mg/l; and are oftennegatively affected at 4.25 mg/l(Davis 1975). Low dissolved oxygenimpairs metabolic rate, growth, swim-ming performance, and overall survivalof young salmonids.

Depth

Water depth usually does notprevent migration because adult steel-head normally migrate when streamflows are relatively high. Thompson(1972) wrote that 18 cm is the minimumdepth required for successfulmigration of adult steelhead. Themigration of adult salmonids is morecommonly hindered by excessive watervelocity or obstacles that impede theswimming or jumping of the fish.

Water depth may be important inthe selection of redd sites.Shapovalov and Taft (1954) stated thatsteelhead redds are rarely exposed byfalling stream levels. Bovee (1978)showed that steelhead most commonlyspawn at depths averaging 36 cm(range, 15-61 cm>. The depths ofWashington winter steelhead reddsranged from 12 to 70 cm (Hunter 1973).Carroll (1984) measured water depthsof 12 to 29 cm over steelhead redds ina Klamath River tributary.

Steelhead tend to occupy theshallow riffle areas of streams,particularly during the first year oflife (Hartman 1965). Bovee (1978)presented probability of use curvesshowing that steelhead fry are mostcommonly found in water 8 to 36 cmdeep, and steelhead juveniles areusually located in water 25 to 50 cmdeep. In the Southwest region steel-head streams are annually subjected tolow flow conditions due to theextended summer-fall dry season; thus,pool frequency and depth are impor-tant. In a 2-year study at SingleyCreek, a small coastal stream justsouth of Cape Mendocino, Cross (1975)found that 67%-96% of young-of-the-year steelhead resided in pools. FromJune to September, the riffle surfacearea was reduced 45% while the surfacearea of pools diminished only 26%.Excessive sediment inputs that fillpools can greatly reduce a stream'scapacity to rear steelhead to smoltsize.

Water Movement

Steelhead may encounter watervelocity barriers during upstreammigration, often at falls or culverts.Velocities of 3-4 m/s begin to greatlyhinder the swimming ability ofsteelhead and may retard migration(Reiser and Bjornn 1979). Thompson(1972) outlined methods to calculateminimum and maximum acceptable stream-flows for migrating adult steelhead,for specific stream sections.

Bovee (1978) showed that steel-head spawn in areas with watervelocities of 30-110 cm/s but that thepreferred velocity approximated60 cm/s. Smith (1973) found that thepreferred water velocity for spawningsteelhead in Oregon ranged from 40 to91 cm/s. Steelhead redds were inareas where velocities were 15 to 54cm/s in a small tributary of theKlamath River (Carroll 1984). Sinceswimming performance improves withsize, large adult steelhead can

11

establish redds in faster currentareas of the stream.

Permeability is defined as thecapacity of the gravel to transmitwater. Different spawning gravelstransmit water at different rates.Several studies have demonstrated thatincreased intragravel velocitiesimprove the survival of steelhead eggsand fry before emergence and also thecondition of fry that emerge (Shumway1960; Silver 1960; Coble 1961; Silveret al. 1963; Shumway et al. 1964).McNeil and Ahnell (1964) concludedthat the streambeds of highly produc-tive spawning streams had gravels withhigh permeability (24,000 cm/h) andhad less than 5% (by volume) sand andsilt that passed through a sieve of0.833 mm mesh.

Sediment

Quantitatively, sediment is thegreatest single pollutant in thenation's water (Ritchie 1972). Ander-son (1971) reported that sedimentproduction in northern Californiawatersheds increased markedly as aresult of poor land managementpractices and floods. The steelhead'senvironment can be impaired both bysediment in suspension and by par-ticles deposited as bedload sediment.About 28% of the total spawning areain a once important 16-mile stretch ofthe upper Trinity River has been lostdue to the deposition of sediment(California Resources Agency 1970).Stream channels of northern Californiamarkedly aggraded after large floodevents during the 1960's and 1970's(Lisle 1982). Channel widening andloss of pool-riffle sequence due toaggradation damaged spawning andrearing habitat of steelhead. Thepool-riffle sequence and pool qualityin some northern California streamsstill had not fully recovered by 1980from a 1964 regional flood (Lisle1982).

For rearing juvenile steelhead,deposited sediment reduces the

,carrying capacity of the streamdirectly by reducing available rearinghabitat and indirectly by reducing theproduction of invertebrate food.Bjornn et al. (1977) found significantreductions in numbers of juvenilesteelhead in stream channels whereboulders were imbedded in sediment.Crouse et al. (1981), who devised avisual substrate scoring system basedon particle size and degree of cobbleembedment, reported that fishproduction was significantlycorrelated with substrate score. Theyreported significant decreases in fishproduction in streams where cobbleswere embedded 80%-100% and wheresediment (2.0 mm or less) composed26%-31% (by volume) of the totalsubstrate composition. The authorsconcluded that rearing habitats ofjuvenile salmonids in streams, as wellas spawning gravels requireprotection from excessive quantitiesof fine sediments.

The size of substrate materialhas been related by numerous investi-gators to the standing crops of inver-tebrates (Sprules 1947; Kimble andWesche 1975). Pennak and VanGerpen(1947) found that the number ofbenthic invertebrates decreased in theprogression from rubble to bedrock togravel to sand. Reiser and Bjornn(1979) reported that aquatic insectproduction was highest in substratecomposed largely of coarse gravel(3.2-6.7 cm) and rubble (7.6-30.4 cm).

Suspended sediment occasionallyreaches concentrations high enough todirectly injure steelhead (3,000 ppmor greater) (Cordone and Kelley 1961).Physiological damage includes theadhesion of silt particles to thechorion of Salmonid ova (Cordone andKelley 1961) and the abrasion,thickening, and fusion of gillfilaments (tierbert and Merkens 1961).Sigler et al. (1984) reported thatchronic turbidity in streams duringemergence and rearing of steelheadaffects the numbers and quality of

12

LI. fish produced. Turbid water also took place when turbidities had

affects recreational angling for decreased to 30 Jackson Turbiditysteelhead. A study on the Eel River Units or less; such low levels ofby the California Department of Fish turbidity occurred during onlyand Game during two winter steelhead one-third of the fishing season (Blakeseasons showed that 85% of all fishing and Goodson 1969).

,

k 1 3

LITERATURE CITED

Allen, K.R. 1969. Limitations onproduction in Salmonid populationsin streams. Pages 3-18 in T.G.Northcote, ed. Symposium on-Troutand Salmon in Streams. Univ. ofB. C., Inst. of Fish., Vancouver.

Anderson, H.W. 1971. Relativecontributions of sediment fromsource areas and transport pro-cesses. Pages 55-63 in J. Morris,ed. Symposium, Forest Land Uses andStream Environment. Oregon StateUniversity Press, Corvallis.

Bali, J.M. 1959. Scale analysis ofsteelhead trout. Salmo aairdnerigairdneri Richardson, from various coastal watersheds of Oreqon. M.S.Thesis. Oregon State University,Corvallis. 189 pp.

Barnhart, R.A. 1975. Pacific slopesteelhead trout management. Pages7-11 in W. King, ed. Wild TroutManagement, Symposium. TroutUnlimited, Denver, Colo.

Bell, M.C. 1973. Fisheries handbookof engineering requirements and bio-logical criteria. U.S. Army Corpsof Engineers, Portland, Oregon.Contract No. DACW 57-68-C-0086. 425pp.

Bjornn, T.C. 1978. Survival, pro-duction and yield of trout and chi-nook salmon in the Lemhi River,Idaho. Coll. of Forestry, Wildl.,and Range Sciences, Univ. of Idaho,Moscow. Bull. No. 27. 57 pp.

15

Bjornn, T.C., M.A. Brusven, M.P. Mol-nav, J.H. Milligan, R.A. Klamt, E.Chaco, and C. Schaye. 1977. Trans-port of granitic sediment in streamsand its effects on insects and fish.Univ. Idaho Coop. Fish. Res. UnitBull. 17. Completion Rep. Proj.B-036-1DA. 43 pp.

Blake, R.B., and L.F. Goodson. 1969.Fishing pressure and turbidity inthe upper Eel River basin. Calif.Dep. Fish Game Rep., Region I. 6 pp.(Mimeo).

Bovee, K.D. 1978. Probability of usecriteria for the family Salmonidae.Instream Flow Information Paper No.4. U.S. Fish Wildl. Serv. FWS/OBS-78/07. 80 pp.

Boydstun, L.B. 1977a. California'ssteelhead program. Pages 26-30 inT.J. Hassler, R.R. Van Kirk, eds.Proceedings, Genetic Implications ofSteelhead Management Symposium.Calif. coop. Fish. Res. Unit,Humboldt State University, Arcata.Spec. Rep. 77-l.

Bovdstun, L.B. 1977b. Lower KlamathRiver tagging study. Calif. Dep.Fish Game. Performance Rep.AFS-20-3. 28 pp. (mimeo).

Brown, G.W. 1971. Water temperaturein small streams as influenced byenvironmental factors. Pages 175-181 in J. Morris,Forest Land

ed. Symposium,Uses and Stream

Environment. Oregon State Univer-sity Press, Corvallis.

Burns, J.W. 1971. The carryingcapacity for juvenile salmonids insome northern California streams.Calif. Fish Game 57:44-57.

Bustard, D.R., and D.W. Narver. 1975.Aspects of the winter ecology ofjuvenile coho salmonkisutch) and steelhead troutgairdneri). J. Fish. Res. Board.Can. 32(5):667-680.

California Department of Fish andGame. 1965. Pages 323-679 inCalifornia Fish and Wildlife Plan,Vol. 3, Part B, The ResourcesAgency, Sacramento.

California Department of Fish andGame. 1975. Steelhead rainbowtrout policy. Pages 407-409 in Fishand Game Code, 1983. Sacramento.

California Resources Agency. 1970.Sediment problems in the TrinityRiver near Lewiston. Task ForceRep. to Sec. for Res. Calif. Res.Agency. 32 pp.

Carroll, E.W. 1984. An evaluation ofsteelhead trout and instreamstructures in a Californiaintermittent stream. M.S. Thesis.Humboldt State University, Arcata,Calif. 51 pp.

Chapman, D.W. 1966. Food and spaceas regulators of Salmonidpopulations in streams. Am. Nat.100:345-357.

Chapman, D.W., and T.C. Bjornn. 1969.Distribution of salmonids instreams, with special reference tofood and feeding. Pages 153-176 inT.G. Northcote, ed. Symposium onTrout and Salmon in Streams. Univ.of B. C., Inst. of Fish., Vancouver.

Coble, D.W. 1961. Influence of waterexchange and dissolved oxygen inredds on survival of steelhead troutembryos. Trans. Am. Fish. Soc.90(4):469-474.

Cordone, A.J., and D.W. Kelley.1961. The influences of inorganicsediment on the aquatic life ofstreams. Calif. Fish Game 42(2):189-228.

Cross, P.D. 1975. Early life historyof steelhead trout (Salmo gairdneri)in a small coastal stream. M.S.Thesis. Humboldt State University,Arcata, Calif. 44 pp.

Crouse, M.R., C.A. Callahan, K.W.Malveg, and S.E. Dominquer. 1981.Effects of fine sediments on growthof juvenile coho salmon laboratory streams. Trans. Am.Fish. Soc. 110(2):281-286.

Davis, G.E., J. Foster, C.E. Warren,and P. Doudoroff. 1963. Theinfluence of oxygen concentration onthe swimming performance of juvenilePacific salmon at varioustemperatures. Trans. Am. Fish. Soc.92(2):111-124.

Davis, J.C. 1975. Minimal dissolvedoxygen requirements of aquatic lifewith emphasis on Canadian species:a review. J. Fish. Res. Board. Can.32(12):2295-2332.

Edmundson, E., F.H. Everest, and D.W.Chapman. 1968. Permanence ofstation in juvenile chinook salmonand steelhead trout. J. Fish. Res.Board. Can. 25(7):1453-1464.

Embody, G.C. 1934. Relation oftemperature to the incubationperiods of eggs of four species oftrout. Trans. Am. Fish. Soc.64: 281-292.

Everest, F.H. 1973. Ecology andmanagement of summer steelhead inthe Rogue River. Oregon State GameComm. Fish. Res. Rep. No. 7,Corvallis. 48 pp.

Everest, F.H., and D.W. Chapman.1972. Habitat selection and spatialinteraction by juvenile chinooksalmon and steelhead trout in two

Idaho streams. J. Fish. Res. Board.Can. 22:1035-1081.

Fite, K.R. 1973. Feeding overlapbetween roach and juvenile steelheadin the Eel River. M.S. Thesis,Humboldt State University, Arcata,Calif. 38 pp.

Forsgren, H.L., II. 1979. Age,growth, origin and repeat spawningof winter steelhead (Salmogairdneri) in Mad River, California.M.S. Thesis. Humboldt State Univer-sity, Arcata, Calif. 56 pp.

Fraser, F.J. 1969. Population den-sity effects on survival and growthof juvenile coho salmon and steel-head trout in experimental streamchannels. Pages 253-266 in T.G.Northcote, ed. Symposium on Troutand Salmon in Streams. Univ. ofB.C., Inst. of Fish., Vancouver.

Fry, D.H., Jr. 1973. Anadromousfishes of California. Calif. Dep.Fish Game, Sacramento. 111 pp.

Hallock, R.J., W.F. VanWoert, and L.Shapovalov. 1961. An evaluation ofstocking hatchery-reared steelheadrainbow- trout (Salmo gairdnerigairdneri) in the Sacramento Riversystem. Calif. Dep. Fish Game FishBull. No. 114. 74 pp.

Hartman, G.F. 1965. The role ofbehavior in the ecology andinteraction of underyearling cohosalmon (Oncorhynchus kisutch) andsteelhead gairdneri).trout (salmoJ. Fish. Res. Board. Can. 22(4):1035-1081.

Herbert, D.W.M., and J.C. Merkens.1961. The effect of suspendedmineral solids on the survival oftrout. Int. J. Air Water Pollut.5(1):46-53.

Hiss, J.M. 1984. Diet of age-0steelhead trout and speckled dace inWillow Creek, Humboldt County,

California. M.S. Thesis, HumboldtState University, Arcata, Calif. 48pp.

Hooper, D.R. 1973. Evaluation of theeffects of flows on trout streamecology. Dep. of Eng. Res., PacificGas and Electric Co., Emeryville,Calif. 97 pp.

Hunter, J.W. 1973. A discussion ofgame fish in the State of Washingtonas related to water requirements.Rep. by Wash. State Dep. Game, Fish.Manage. Div., Wash. State Dep. Ecol.66 pp.

Jensen, P.T. 1971. Salmon andsteelhead. Pages 1-13 in Report tothe State Water Resources ControlBoard. Calif. Fish Game Environ.Serv. Admin. Rep. No. 71-2.

Jones, D.E. 1974. The study ofcutthroat-steelhead in Alaska.Alaska Dep. Fish Game, Div. of SportFish. Anadromous Fish Studies.Annu. Prog. Rep. 1973-1974, StudyAFS-42-2. 31 pp.

Kerstetter, T.H., and M. Keeler.1976. Smolting in steelhead troutSalmostudy of populations two different

g a i r d n e r i : at a comparative

hatcheries and the Trinity River,northern California, using gill Na,K, ATPase assays. Humboldt StateUniversity Sea Grant Project,Arcata, Calif. HSU-SG9. 26 pp.

Kesner, W.D., and R.A. Barnhart.1972. Characteristics of the fall-run steelhead trout (Salmo aairdnerigairdneri) of the Klamath Riversystem with emphasis on the half-pounder. Calif. Fish Game58(3):204-220.

Kimble, L.A., and T.A. Wesche. 1975.Relationship between selectedphysical parameters and benthiccommunity structure in a smallmountain stream. Univ. Wyo.,Laramie. Water Resour. Res. Inst.Series 55. 64 pp.

17

Kralik, N.J., and J.E. Sowerwine.1977. The role of two northernCalifornia intermittent streams inthe life history of anadromoussalmonids. M.S. Thesis. HumboldtState University, Arcata, Calif. 68pp.

Kreb, M. 1984. California Conser-vation Corps project 201. Pages lo-ll in K. Hashagen, C. Toole, B.Wyatt, S. Sommarstrom, S. Taylor,eds. Report of the 2nd Calif.Salmon and Steelhead RestorationConference. Univ. Calif. Sea Grant,UCSG-MAP-21, Davis. 97 pp.

Lantz, R.L. 1971. Influence of watertemperature on fish survival, growthand behavior. Pages 182-193 in J.Morris, ed. Symposium, Forest LandUses and Stream Environment. OregonState University, Corvallis. 252pp.

LeBrasseur, R . J . 1966. Stomachcontents of salmon and steelheadtrout in the northeastern PacificOcean. J. Fish. Res. Board. Can.23(1):85-100.

Lisle, T.E. 1982. The recovery ofstream channels in north coastalCalifornia from recent large floods.Pages 31-41 in K. Hashagen, ed.Habitat Disturbance and RecoveryProceedings. Calif.Francisco. 90 pp.

Trout, San

Manzer, J.I. 1968. Food of Pacificsalmon and steelhead trout in thePacific Ocean. J. Fish. Res. Board.Can. 25(5):1085-1089.

McConnell, R.J., and G.R. Snyder.1972. Key to field identificationof anadromous juvenile salmonids inthe Pacific northwest. NOAA (Natl.Ocean. Atmos. Adm.) Tech. Rep. NMFS(Natl. Mar. Fish. Serv.) Circ. 366.6 pp.

McFadden, J.T. 1969. Dynamics andregulation of Salmonid populationsin streams. Pages 313-329 in T.G.

Northcote, ed. Symposium on Salmonand Trout in Streams. Univ. B. C.,Inst. Fish., Vancouver.

McNeil, W.J., and W.H. Ahnell. 1964.Success of pink salmon spawningrelative to size of spawning bedmaterials. U.S. Fish Wildl. Serv.Spec. Sci. Rep. Fish. No. 469. 15pp.

Meigs, R.C., and C.P. Pautzke. 1941.Additional studies on the life his-tory of the Puget Sound steelhead(Salmo gairdneri). State Wash. Dep.Game, Bio.Bull.. No. 5. 13 pp.

Menchen, R.S. 1981. Predation byyearling steelhead Salmo gairdneri,released from Coleman National FishHatchery, on naturally produced chi-nook salmon Oncorhynchus tshawytschafry and eggs in Battle Creek, 1975.Calif. Dep. Fish Game, AnadromousFisheries Branch Office Report,Sacramento. 6 pp.

Milne, D.J. 1948. The growth,morphology and relationship of thespecies of Pacific salmon and thesteelhead trout. Ph.D. Thesis.McGill University, Montreal. 101 pp.

Moore, M.R. 1980. Factors influ-encing the survival of juvenilesteelhead rainbow trout. Salmogairdneri in the Ventura River,California. M.S. Thesis. HumboldtState University, Arcata, Calif. 82pp.

Moyle, P.B. 1976. Inland fishes ofCalifornia. University of Cali-fornia Press, Berkeley. 405 pp.

National Council on Gene Resources.1982. Anadromous Salmonid geneticresources: an assessment and planfor California. California GeneResource Program, Berkeley. 168 pp.

Needham, P.R., and R. Gard. 1959.Rainbow trout in Mexico and Cali-fornia with notes on the cutthroat

18

L series. Univ. Calif. Publ. Zool.67:1-124.

Overton,m K. 1984. U.S. ForestService Six Rivers National Forest.Page 37 in K. Hashagen, C. Toole, B.Wyatt, S. Sommarstrom, S. Taylor,eds. Report of the 2nd Calif.Salmon and Steelhead RestorationConference. Univ. Calif. Sea Grant,UCSG-MAP-21, Davis. 97 pp.

Overton K., W. Brock, J. Moreau, andJ. Boberg. 1981. Restoration andenhancement program of anadromousfish habitat and populations on SixRivers National Forest. Pages 158-168 in T.J. Hassler, ed. Proceed-ings:- Propagation, Enhancement, andRehabilitation of Anadromous Sal-monid Populations and HabitatSymposium. Humboldt State Univer-sity, Arcata, Calif. 168 pp.

Pennak, R.W., and E.D. VanGerpen.1947. Bottom fauna production and

L

physical nature of the substrate ina northern Colorado trout stream.Ecology 28:42-48.

Peterson, G.R. 1966. The relation-ship of invertebrate drift abundanceto the standing crop of benthicorganisms in a small stream. M.S.Thesis. University of BritishColumbia, Vancouver. 39 pp.

Phillips, R.W., and H.J. Campbell.1962. The embryonic survival ofcoho salmon and steelhead trout asinfluenced by some environmentalconditions in gravel beds. Pac.Mar. Fish. Comm. 14th. Annu. Rep.1961: 60-75.

Pollard, H.A., II, and T.C. Bjornn.1973. The effects of angling andhatchery trout on the abundance ofjuvenile steelhead trout. Trans.Am. Fish. Soc. 102(4):745-752.

L

Ponce, S.L. 1974. The biochemicaloxygen demand of finely dividedlogging debris in stream water.Water Resour. Res. 10(5):983-988.

Rawstron, R., and K. Hashagen. 1984.California Department of Fish andGame salmonid enhancement andrestoration activities. Page 12 inK. Hashagen, C. Toole, B. Wyatt, S.Sommarstrom, S. Taylor, eds. Reportof the 2nd Calif. Salmon andSteelhead Restoration Conference.Univ. Calif. Sea Grant, USCG-MAP-21,Davis. 97 pp.

Reeves, G.H. 1979. Population dynam-ics of juvenile steelhead in rela-tion to density and habitat charac-teristics. M.S. Thesis. HumboldtState University, Arcata, Calif. 67pp.

Reingold, M. 1968. Water temperatureaffects the ripening of adult fallchinook salmon and steelhead. Prog.Fish-Cult. 30(1):41-42.

Reiser, D.W., and T.C. Bjornn. 1979.Habitat requirements of anadromoussalmonids. 54 pp. in W.R. Meehan,ed. Influence of Forest and RangeManagement on Anadromous FishHabitat in Western North America.Pacific N.W. Forest and Range Exp.Sta. USDA For. Serv., Portland.Gen. Tech. Rep. PNW-96.

Ritchie, J.C. 1972. Sediment, fishand fish habitat. J. Soil WaterConser. 27(3):124-125.

Roelofs, T.D. 1983. Current statusof California summer steelhead(Salmo gairdneri) stocks and habitatand recommendations for theirmanagement. USDA For. Serv., Region5, San Francisco, Calif. 77 pp.

Royal, L.A. 1972. An examination ofthe anadromous trout program of theWashington State Game Department.Wash. State Dep. Game, Final Rep.AFS-49, Olympia. 176 pp.

Shapovalov, L., and A.C. Taft. 1954.The life histories of the steelheadrainbow trout (Salmo

gairdnerigairdneri) and silver salmon(Oncorhynchus kisutch) with special

19

reference to Waddell Creek, Cali-fornia, and recommendations regard-ing their management. Calif. Dep.Fish Game Fish. Bull. 98. 375 pp.

Sheppard, D. 1972. The presentstatus of the steelhead trout stocksalong the Pacific coast. Pages 519-556 in D.H. Rosenberg, ed. A Reviewof the Oceanography and RenewableResources of the Northern Gulf ofAlaska. IMS Rep. R72-23, Sea GrantRep. 73-3. Institute of MarineScience, University of Alaska,Fairbanks. 690 pp.

Shumway, D.L. 1960. The influence ofwater velocity on the development ofSalmonid embryos at low oxygenlevels. M.S. Thesis. Oregon StateUniversity, Corvallis. 49 pp.

Shumway, D.L., C.E. Warren, and P.Doudoroff. 1964. Influence ofoxygen concentration and watermovement on the growth of steelheadtrout and coho salmon embryos.Trans. Am. Fish. Soc. 93(4):342-356.

Sigler, J.W., T.C. Bjornn, and F.H.Everest. 1984. Effects of chronicturbidity on density and growth ofsteelheads and coho salmon. Trans.Am. Fish. Soc. 113:142-150.

Silver, S.T. 1960. The influence ofwater velocity and dissolved oxygenon the development of Salmonidembryos. M.S. Thesis. Oregon StateUniversity, Corvallis. 50 pp.

Silver, S.J., C.E. Warren, and P.Doudoroff. 1963. Dissolved oxygenrequirements of developing steelheadtrout and chinook salmon embryos atdifferent water velocities. Trans.Am. Fish. Soc. 92(4):327-343.

Smith, A.K. 1973. Development andapplication of spawning velocity anddepth criteria for Oregon salmonids.Trans. Am. Fish. Soc. 102:312-316.

Smith, S.B. 1960. Racial charac-teristics in stocks of anadromous

rainbow trout, Salmo gairdneriRichardson. Ph.D. Thesis. Univer-sity of Alberta, Edmonton, Alberta.

Smith, S.B. 1969. Reproductiveisolation in summer and winter racesof steelhead trout. Pages 21-38 inT.G. Northcote, ed. Symposium onSalmon and Trout in Streams. Univ.B. C., Inst. Fish. Vancouver.

Snyder, J.O. 1925. The half-pounderof Eel River, a steelhead trout.Calif. Fish Game 11(2):49-55.

Sprules, W.M. 1947. An ecologicalinvestigation of stream insects inAlgonquin Park, Ontario. Univ.Toronto Stud. Biol. 56., Publ. Ont.Fish. Res. Lab. 69:1-81.

Swift, C.C. 1975. Survey of thefreshwater fishes and their habitatsin the coastal drainages of southernCalifornia. Natural History Museumof Los Angeles County, California.364 pp.

Tebo, L.B., Jr. 1974. Review ofselected parameters of trout streamquality. Pages 20-32 in Symposium

Trout Habitat Research andManagement Proceedings. Appala-chian Consortium Press, Boone, N.C.

Thompson, K. 1972. Determiningstream flows for fish life. Pages31-50 in Proceedings, Instream FlowRequirement Workshop. Pac. N.W.River Basin Comm., Vancouver, Wash.

Wagner, H.H. 1967. A summary ofinvestigations of the use ofhatchery-reared steelhead in themanagement of a sport fishery.Oregon State Game Comm. Fish. Rep.5. 62 pp.

Wagner, H.H. 1974. Photoperiod andtemperature regulation of smoltingin steelhead trout (Salmo gaird-neri). Can. J. Zool. 52:805-8 12.

20

Wales, J.H. 1941. Development ofsteelhead trout eggs. Calif. FishGame 27(4):250-260.

Wesche, T.A., and P.A. Rechard. 1980.A summary of instream flow methodsfor fisheries and related researchneeds. Univ. Wyo., Water Resour.Res. Inst., Eisenhower ConsortiumBull. 9. 122 pp.

Wickett, W.P. 1954. The oxygensupply to salmon eggs in spawningbeds. J. Fish. Res. Board. Can.11(6):933-953.

Withler, I.L. 1966. Variability inlife history characteristics ofsteelhead trout (Salmo gairdneri)along the Pacific coast of NorthAmerica. J. Fish. Res. Board. Can.23(3):365-393.

21

REPORT D O C U M E N T A T I O N 1. REPORT No- 3 . RECIPIENT'S Accession N o

PAGE Biological Report 82(11.60)*1**. Title and Subhtle Species Profiles: Life tiistories and Environmental 5. Ww?l oat*

Requirements of Coastal Fishes and Invertebrates (Pacific South- June 1986west) -- Steelhead b.

. AlJthortS)

Roger A. Barnhart Performing Organization Name and Address

California Cooperative Fishery Research UnitHumboldt State UniversityArcata, CA 95521

2. Sponsoring Organization N a and Address

9. Performing Organizaion Rept. N o

10. l ro~~ctflarh/Work Unct No.

11. COnWrctfC) or GranUG) No.

(C)

(G)

National Coastal Ecosystems Team U.S. Army Corps of Engineers 11. 'lyCUOfRW#Or( & r.nOd Cownd

Fish and Wildlife Service Waterways Experiment StationU.S. Department of the Interior P.O. Box 631Washington, D.C. 20240 Vicksburg, MS 39130

m

14.

IS. Supphmontary Notas

* U.S. Army Corps of Engineers, TR EL-82-46 . Abstract (Limit: 200 words)

Species profiles are summaries of the literature on the taxonomy, life history, andenvironmental requirements of coastal fishes and aquatic invertebrates. They areprepared to assist with environmental impact assessment. The steelhead Salmo gairdneri,an anadromous rainbow trout, supports an important sport fishery in the PacificSouthwest. Although native populations of steelhead have declined, these fishannually enter coastal streams from northern to southern California in years whenwinter stream flow is high. Steelhead ascend coastal streams from the ocean tospawn in cool, well-oxygenated waters with suitable depth, current velocity, andgravel size. After hatching, steelhead fry emerge from the gravel and begin a fresh-water rearing phase that generally extends from 1 to 3 years. Rearing habitat withproper environmental conditions is extremely important to steelhead production.Excessive sedimentation reduces food production, pool depth, and cover--all importantto juvenile steelhead survival. Steelhead smolts migrate during spring to saltwater,where most of their growth and sexual maturity is attained in 1 or 2 years. Attemptshave increased to protect wild steelhead stocks, to maintain existing spawning andrearing habitat, to restore or enhance degraded habitat where feasible, to useartificial propagation efficiently, and to establish fishing regulations that providequality angling for steelhead.

17 . Document Analysis Descriptors

Trout SalinityStreams TemperatureFisheries Sediments

Feeding habits GrowthFood chains OxygenLife cycles Animal migrations

b . Identifiers/Open-Ended Terms

Steelhead troutSalmo gairdneriSpawning requirementsRearing requirementsc. C o s a t i Field/Group

18. AvarIabIlity Sta(oment

Unlimited

lg. kcurlty Class CTh,s Rcoon)

I -

21. No. Of PJWS

Unclassified 21---to. Socur~ty CIass CThls Paw)

Iz. Pncc

Unclassified(See ANSI-Z19.18) “r I ,“I.__ . _.*- _. - ,

(Formerly NTIS-35)Decwtment of Commerce